Andesianidae, a New Family of Monotrysian Moths (Lepidopteraiandesianoidea) from Austral South America
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CSIRO PUBLISHING www.publish.csiro.au/journals/is Invertebrate Systematic.?, 2003, 17, 15-26 Andesianidae, a new family of monotrysian moths (LepidopteraiAndesianoidea) from austral South America Donald R. Davis ' and Patricia Gentili ^Department of Systematic Biology-Entomology, National Museum of Natural History, PO Box 37012, Smithsonian Institution, Washington, DC 20560-0127, USA. BTo whom correspondence should be addressed. Email: [email protected] Abstract. The new family Andesianidae is proposed for the genus Andesiana, consisting of three previously described species, A. brunnea Gentili, A. lamellata Gentili and A. similis Gentili, from southern Argentina and Chile. Originally described in the ditrysian family Cossidae, Andesiana has been found to possess a monotrysian female reproductive system. The principal autapomorphies for the family are the elongate second segment of the labial palpi, presence of a male tibial pouch and associated femoral hair pencil and, within the monotrysian Heteroneura, possibly the strongly dimorphic antennae (broadly bipectinate in male, filiform in female). Other significant characters of the group include a greatly reduced haustellum, male gen i tali a with a broad vinculum, acuminate uncus, scutiform juxta and a pronounced saccular lobe of the valva that bears a subapical, setose appendage within a shallow, mesal cavity. The female genitalia possess a short cloaca and paired anal papillae and lack signa. Although known to be retained only in the female of A. lamellata, the presence of a two-branched Rs in the hindwing may indicate an origin of the Andesianidae basal to the Nepticuloidea. In contrast, the structure of the wing coupling apparatus for the family suggests a later appearance. Because no synapomorphy is known linking the Andesianidae with any other superfamily, the new superfamily, Andesianoidea, is proposed. Introduction The monotrysian Heteroneura, a questionably mono- Field investigations conducted in the southern temperate phyletic assemblage of five distantly related superfamities, is regions of South America, largely during the decade of believed to represent the basal group of the infraorder 1979-1989, significantly increased our knowledge of the Heteroneura, as indicated by the (1) retention of a primitive more primitive, basal groups of Lepidoptera. As a direct monotrysian reproductive system (Dugdale 1974), with a result of these collective efforts, three new families of moths single reproductive opening in the female, ventral common have been documented including the Heterobathmiidae oviduct and absence of a tubular ductus seminalis; (2) (Kristensen and Nielsen 1979, 1983), Palaephatidae (Davis presence of a tergosternal connection (Kyrki 1983) 1986) and, in this report, Andesianidae. Another result of immediately caudad to the first abdominal spiracle (Fig. 23), that fieldwork was the discovery of four families previously formed by a ventrocaudal process from tergum I connecting not known to occur in either the New World (Neopseustidae, with the anterolateral extension of sternum II; and (3) often Davis 1975 and Davis and Nielsen 1980; Eriocottidae, Davis more generalised distribution of wing microtrichia. Major 1990), South America (Micropterigidae, Kristensen and synapomorphies shared with the ditrysian Heteroneura Nielsen 1982), or the southern hemisphere (Prodoxidae, include: (/) reduction of the hindwing Rs system (i.e. Nielsen and Davis 1985). heteroneurous venation, with Sc and R fused from the basal The genus Andesiana Gentili, with three included species fourth of the wing to the wing margin and Rs typically {A. brunnea Gentili, A. lamellata Gentili and A. similis unbranched); (2) frenulum-retinaculum wing coupling; (3) Gentili) from southern Argentina and Chile was proposed composite male frenulum; (4) presence of precoxal bridge originally in the ditrysian family Cossidae (Gentili 1989). (Nielsen and Kristensen 1996); and (5) loss of the first Later questioning the family placement of this genus, PG abdominal sternum (Brock 1971; Kristensen 1984). then requested DRD to examine specimens of Andesiana. Monophyly of the monotrysian Heteroneura (or Monotrysia) After dissecting a female, DRD recognised that it is inadequately supported, with two possible synapo- represented a new family of monotrysian moths. morphies proposed (Davis 1999): (7) presence of spine B CSIRO 2003 I0.107I/IS02006 1445-5226/03/010015 16 Invertebrate Systematic,1* D. R. Davis and P. Gentili .4,\ %%%'% . V ' -if-. & • k Figs 1-4. Adults of Andesiana. 1, Andesiana lamellata, male (45 mm); 2, Andesiana lamellata, male (45 mm); 3, Andesiana lamellata, female (54 mm); 4, Andesiana similis, male (35 mm). (Wingspan in parentheses). combs (pectinifers) on the male valva (absent in whose larvae are known to be stem borers. Consequently, Tischerioidea, Palaephatoidea and Andesiana); and (2) adult Andesiana may be easily mistaken for a member of the complete or partial subdivision of the second abdominal ditrysian family Cossidae, unless the female internal sternum into a smaller anterior sclerite (S2a) and a usually reproductive morphology is examined. Although their much larger caudal sclerite. The latter condition is often biology is unknown, the greasy condition, which develops in associated with the development of a pair of oval areas many museum specimens, as well as their relative body size, (windows) that largely separate the more sclerotised strongly suggests that larval Andesiana are also stem borers. anterior/posterior regions of sternum II. Lightly sclerotised Adult Andesianidae are phototrophic and are readily windows also occur on S2 in some ditrysian families (e.g. attracted to ultraviolet light. Acrolophidae). Partial to complete subdivision of sternum II, Andesianidae is characterised by at least two often associated with paired sternal windows, is known to autapomorphies: (7) elongate second segment of the labial exist in other primitive glossatan families (e.g. Eriocraniidae palpi (Fig. 5) and (2) presence of male tibial pouch and and Acanthopteroctetidae, Davis 1978; Lophocoronidae, associated femoral hair pencil (Fig. la, b). The second labial Nielsen and Kristensen 1996) and may be part of the palpal segment is nearly 4 % the length of the apical segment. Lepidoptera Glossata groundplan. The latter terminates in an elongate sensory pit (organ of Adult Andesiana are moderately large, heavy bodied, vom Rath; Figs 5, 14), which often is found everted in dried cossoid-appearing moths quite unlike any previously specimens (Fig. 12). Specialised organs for pheromone recognised member of the Monotrysia Heteroneura. Their dispersion are sparingly developed among the monotrysian wing span ranges from 27 to 61 mm (1.19-2.38 inches), with Heteroneura, with the notable exception of Palaephatidae a maximum size nearly twice that of the largest previously (Davis 1986) and one unusual occurrence within the known monotrysian species (e.g. 32 mm in female Incurvarioidea (Wojtusiak 1999). The males of Andesiana Palaephatus dimorphus Davis). Moreover, the bodies and possess a rather elaborate androconial organ on the hind leg wings of dried specimens often become greasy, a condition consisting of an elongate hair pencil composed of numerous typically occurring in other families, particularly Cossidae, filamentous setae that arise subapically from the femur and The new monotrysian moth family Andesianidae Invertebrate Systematics 17 hf \ S= R1 Rs1 Figs 5-7. Adult morphology of Andesiana. Andesiana lamellatu 5-7a: 5, frontal view of head (1.0 mm); 6, female wing venation; 7, male legs (5.0 mm); 7a, detail of male femoral hair pencil and tibial pouch (0.5 mm). 7b, Andesiana similis: male femoral hair pencil and inflated tibial pouch (1.0 mm). (HP = hair pencil; H - haustellum; IS = intercalary sclerite; OR = organ of vom Rath; TP = tibial pouch; scale lengths in parentheses). extend approximately 1/2-2/3 the length of the tibia. The external wall of the pouch is densely covered with minute, distal fourth to one third of the hair pencil normally rests non-deciduous, piliform scales. Hydrostatic pressure within inside an inverted pocket within a large, membranous, the pouch everts the pocket, thereby expelling the hair pencil. inflatable pouch at the base of the hind tibia (Fig. la). The Once expelled, it is unlikely that the hair pencil can ever be 18 Invertebrate Systematics D. R. Davis and P. Gentili Figs 8-13. Andesiana lamellata, head morphology. 8, Female antenna, near basal third of flagellum, top of photo = dorsal. 9, Detail of sensillum coeloconicum (centre) from Fig. 8. 10, Male antenna! pectinations, dorsal view. 11, Venter of Fig. 10. 12, Labial palpal segment III with subapical organ of vom Rath exerted. 13, Detail of apical sensillac in Fig. 12. reinserted into the tibial sac. The length of the fully inflated dense scattering of long sensillae chaetica (Fig. 11), which pouch ranges from approximately half the length of the tibia are more dispersed over the female antenna (Fig. 8). in A. lamellata to nearly its entire length in A. similis (Fig. Sensillae coeloconica are scattered along the lower surfaces lb). No males of A. brunnea were available for study. of the male rami and are denser ventrally along the female Strongly dimorphic antennae (broadly bipectinate in flagellomeres (Figs 8, 9). A broadly triangular intercalary male, filiform in female) (Figs 8, 10, 11) may constitute sclerite is present between the posterioventral margins of the another autapomorphy for Andesiana.